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752-4001-00L Mikrobiologie

Julia Vorholt
Lecture 9:
Phototrophy and autotrophy
Global carbon and nitrogen cycles
Nov 19, 2012

Brock Biology of Microorganisms, Twelfth Edition


Madigan / Martinko / Dunlap / Clark
Copyright 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

1)
2)
3)
4)
5)
6)
7)

Nutrients and microbial growth


Introduction to principles of metabolism
Chemoorganotrophy
Chemolithotrophy
Phototrophy
Autotrophy, nitrogen fixation
Global carbon, nitrogen, sulfur cycles

Brock Biology of Microorganisms, Twelfth Edition


Madigan / Martinko / Dunlap / Clark
Copyright 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Photophosphorylation and ETP

ELQ = h c NA -1 = h v
ELQ, Free energy of light quanta
h, Planck constant (6.6 x 10-34 J s)
c, the speed of light (3 x 108 m s-1)
NA, Avogadros number (6 x 1023)
v, frequency

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Types of Photosynthesis

Phototrophs
Purple and green bacteria

Cyanobacteria, algae, green plants

Oxygenic

Anoxygenic
Reducing power

Carbon

electrons

Energy

Reducing power

Carbon

Light

Energy

Light

PS I and II

PS I or II

Van Niel, 1930: Photosynthesis


CO2 + 2 H2A -> [CH2O] + 2 A + H2O
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13.1 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 13.2

Groups of Phototrophs

Green
Sulfurbacteria
Green
Nonsulfurbacteria
Cyanobacteria

Heliobacteria

(Gram +)

Purple bacteria (Proteobacteria)

Purple Nonsulfur Bacteria

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Purple Sulfur Bacteria

Evolutionary Aspects
Eon

BYA Organisms,
events
0

Sunlight represents the most


important energy source on earth.
Phototrophs use light as sole
energy source.
They are the starting point of the
food chain.
The oxygenic photosynthesis
evolved rather early in evolution,
2.7 billion years ago.
=> major consequences in terms of
primary biomass production
(unlimited availability of H2O) and
the presence of O2.

Phanaerozoic

O2
level

Extinction of the
dinosaurs

0.5

Early animals

1.0

Multicellular
eukaryotes

Metabolic
highlights

Cambrian

Precambrian

20%
10%

Proterozoic
1.5

2.0

First eukaryotes
with organelles
Ozone shield

1%

2.5

Great oxidation
event

0.1%

Endosymbiosis?
Aerobic respiration
Oxygenic photosynthesis
(2H2O O2 4H)

Cyanobacteria

3.0

Archaean

Sulfate reduction
Fe3 reduction
3.5

Hadean

Purple and green


bacteria

Anoxygenic photosynthesis

Anoxic

Bacteria/Archaea
divergence

Acetogenesis

4.0

First cellular life; LUCA


Formation of
crust and oceans

Methanogenesis

4.5

Formation of Earth

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Sterile Earth

Fig. 16.6

Chlorophylls and Bacteriochlorophylls


Organisms must produce some form of chlorophyll (or bacteriochlorophyll) to be
photosynthetic
Chlorophyll is a porphyrin
They are part of the reaction center (participate directly in the conversion of light
energy to ATP)
Number of different types of chlorophyll exist with different absorption spectra

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Fig. 13.3

Carotenoids and Phycobilins


Phototrophic organisms have accessory pigments in
addition to chlorophyll, including carotenoids
Carotenoids
Always found in phototrophic organisms
Typically yellow, red, brown, or green
Energy absorbed by carotenoids can be transferred to a
reaction center
Prevent photo-oxidative damage to cells

-carotene
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Fig. 13.8

Anoxygenic Photosynthesis
Anoxygenic photosynthesis is
found in at least four phyla of
Bacteria
Electron transport reactions
occur in the reaction center of
anoxygenic phototrophs
Reducing power for CO2 fixation
comes from reductants present
in the environment (i.e., H2S,
Fe2+, or NO2-)

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Fig. 17.2
Fig. 17.6

Arrangement of Light-Harvesting Chlorophylls


RC, reaction center
LH, light harvesting molecules

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13.2 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Fig. 13.6

Structure of Reaction Center in Purple Bacteria

Arrangement of Pigment Molecules


in Reaction Center

Molecular Model of the Protein Structure


of the Reaction Center

1988, Nobel prize for the structure of the reaction center of Rhodopseudomonas viridis:
J. Deisenhofer, H. Michel, and R. Huber
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Fig. 13.13

Electron Flow in Anoxygenic PS in a Purple Bacterium


1.0
Strong
electron
donor

0.75

0.5

E0
(V)0.25

Cyclic electron
flow (generates
proton motive
force)

0.0
External electron
donors (H2S, S2O32-,
S0, Fe2+)

0.25
Poor
electron
donor

0.5
Red or infrared light
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as Pearson Benjamin Cummings
Chap. 13.4

Fig. 13.14

Arrangement of Protein Complexes in Reaction Center

Light

Out

(periplasm)

Quinone
pool

ATPase

Photosynthetic
membrane

In

(cytoplasm)

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Fig. 13.15

Electron Flow in Purple, Green, Sulfur and Heliobacteria

Purple bacteria

Green sulfur bacteria

Heliobacteria

1.25
1.0
0.75
0.5

E0 (V)

0.25

Reverse
electron
flow

0.25

0.5
Light

Light

Light

PSII

PSI

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PSI

Fig. 13.17

Electron Flow in Oxygenic Photosynthesis


1.25

The Z Scheme:
1.0

PSII PSI

0.75

Cyclic electron
flow (generates
proton motive
force)

0.5

0.25

E0
0.0
(V)
0.25

Noncyclic
electron flow
(generates
proton motive
force)
Light

0.5

Photosystem I

0.75
1.0

Photosystem II

Light

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Fig. 13.18

Oxygenic Photosynthesis
Oxygenic phototrophs use light to generate ATP and
NADPH
The two light reactions are called photosystem I and
photosystem II
Z scheme of photosynthesis
Photosystem II transfers energy to photosystem I

ATP can also be produced by cyclic


photophosphorylation

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Water blooms in lakes consisting of Cyanobacteria

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Autotrophic CO2-fixation

CO2 + 4 [H] + n ATP -> (CH2O) + H2O + n ADP + n Pi

6 different CO2 fixation pathways are known:


1.
2.
3.
4.
5.
6.

Calvin-Benson-Bassham cycle
Reductive citric acid cycle
Reductive acetyl-CoA pathway
3-Hydroxypropionate/malyl-CoA cycle
3-Hydroxypropionate/4-hydroxybutyrate cycle
Dicarboxylate/4-hydroxybutyrate cycle

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Chap.
13.12-13

The Calvin Cycle

12 3-PhosphoRubisCO

glycerate
(36 carbons)

6 Ribulose
1,5-bisphosphate
(30 carbons)

12 1,3-Bisphosphoglycerate
(36 carbons)

Phosphoribulokinase

Fixes CO2 into cellular


material for autotrophic
growth
Requires NADPH, ATP,
ribulose bisphophate
carboxylase (RubisCO),
and phosphoribulokinase
6 molecules of CO2 are
required to generate one
molecule of glucose

12 Glyceraldehyde
3-phosphate
(36 carbons)

6 Ribulose
5-phosphate
(30 carbons)

Sugar
rearrangements

10 Glyceraldehyde
3-phosphate
(30 carbons)

Fructose
6-phosphate
(6 carbons)

To biosynthesis

Overall stoichiometry:
6 CO2 12 NADPH 18 ATP
12 NADP 18 ADP 17 Pi

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Chap.
13.12

C6 H12 O6(PO3 H2)

Fig. 13.30

(Oxidized)

Nitrogenase and Nitrogen Fixation

(Reduced)

Only certain prokaryotes can fix nitrogen


Some nitrogen fixers are free living and others are symbiotic
Products
Nitrogenase
substrates
Reaction is catalyzed by nitrogenase
Sensitive to the presence of oxygen

A wide variety of nitrogenases use different metal cofactors,


usually molybdenum

Overall reaction

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Chap.
13.14

Symbiotic Nitrogen Fixation

Soy bean

Root nodules

N2

NH3

with
Bradyrhizobium
(Alphaproteobacterium)

without

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(Fig. 25.7)
Fig. 25.8

Symbiotic Nitrogen Fixation

The mutalistic relationship between


leguminous plants and nitrogen-fixing
bacteria is one of the most important
symbioses known
Examples of legumes include soybeans,
clover, alfalfa, beans, and peas
Rhizobia are the most well-known
nitrogen-fixing bacteria engaging in
these symbioses

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Key terms Metabolism - I

We need to distinguish between energy sources and carbon sources


(both are usually identical for chemoorganotrophic organisms)
Chemotrophy (energy from chemical substrates)
> Chemoorganotrophy (organic substrates)
> Chemolithotrophy (inorganic substrates)

Energy
sources

Phototrophy (energy from light)


Autotrophy (CO2 as main carbon source)
Heterotrophy (organic compounds as main carbon source)

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Carbon
sources

Key terms Metabolism - II

Aerobic: Growth of an organism using oxygen as electron acceptor


Anaerobic: Growth of an organism not using oxygen as electron
acceptor
Aerotolerant: Anaerobic organism whose growth is not inhibited by
oxygen and does not use it as electron acceptor
Oxic: Environmental condition where oxygen is present
Anoxic: Environmental condition without oxygen

Oxygenic: Type of photosynthesis, oxygen is released from water


Anoxygenic: Type of photosynthesis, without oxygen release

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The Carbon Cycle and Major Carbon Reservoirs on Earth

CO2
Human
activities
Respiration
Land
plants

Animals and
microorganisms
Aquatic
CO2
plants and
phytoplankton Biological pump

Fossil
fuels

Humus
Soil formation

Death and
mineralization
Earths crust

Aquatic
animals

CO2

Rock formation

Carbon is cycled through all of Earths major carbon reservoirs, i.e., atmosphere, land, oceans, sediments,
rocks, and biomass
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Fig. 24.1

Methane hydrate
A methane hydrate is a cage-like
lattice of ice, inside of which are
trapped molecules of methane.

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Pictures by K. Kvenvolden and GEOMAR

The Carbon Cycle

(CH2O)n

Cyanobakterien
Pflanzen

Oxygene
Photosynthese

Tiere,
verschiedene
Mikroorganismen
(Paracoccus, E. coli)

Aerobe Atmung

(Chemolithotrophe)
Aerobe
Methanoxidation

CH4

CO2

Anaerobe
Methanoxidation

Oxisch
Anoxisch

Methanogenese

Anoxygene
Photosynthese

Anaerobe Atmung

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(CH O)

2
n
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24.1 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

E. coli
Paracoccus denitrificans

(Fig. 24.2)

Assimilation/Baustoffwechsel

Anaerober Prozess
Aerober Prozess

Brock Biology of Microorganisms, Twelfth Edition


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Copyright 2009 Pearson Education Inc., publishing as Pearson Benjamin Cummings

Dissimilation/
Energiestoffwechsel

The Carbon Cycle


Phototrophic organisms are the foundation of the carbon cycle
CO2 is fixed primarily by photosynthetic land plants and marine
microbes
CO2 is returned to the atmosphere by respiration of animals and
chemoorganotrophic microbes as well as anthropogenic activities
Microbial decomposition is the largest source of CO2
released to the atmosphere
The carbon and oxygen cycles are intimately linked
Plants dominant phototrophic organisms of terrestrial
environments
The two major end products of decomposition are CH4 and CO2

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The Nitrogen Carbon


Cycle

Rhizobien
Cyanobakterien

N2-Fixierung

Org. N
(NH2-Gruppen

N2

NH3

N2O

Ammonifikation
E. coli

Denitrifikation
Pseudomonas
Paracoccus

NO

Nitrifikation
Nitrosobakterien
(z.B. Nitrosomonas)

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Nitrobakterien
(z.B. Nitrobacter)

NO3-

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(Fig. 24.7)

The Nitrogen Cycle

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Fig. 24.7

The Nitrogen Cycle

N2 is the most stable form of nitrogen and is a major reservoir


The ability to use N2 as a cellular nitrogen source
(nitrogen fixation) is limited to only a few prokaryotes
Denitrification is the reduction of nitrate to gaseous nitrogen
products and is the primary mechanism by which N2 is
produced biologically
Ammonia produced by nitrogen fixation or ammonification can
be assimilated into organic matter or oxidized to nitrate
Denitrification and anammox result in losses of nitrogen from
the biosphere

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The Sulfur Cycle


Org. S

Chemolithotrophe
S-Oxidation

H 2S

Diss. S0
reduktion

Beggiatoa

Photolithotrophe
Dissimilatorische
Sulfatreduktion

Assimilatorische
Sulfatreduktion

Desulfovibrio

S0

SchwefelPurpurbakterien
SchwefelGrne Bakterien

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(Fig. 24.8)

The Sulfur Cycle

Hydrogen sulfide is a major volatile sulfur gas that is produced


by bacteria via sulfate reduction or emitted from geochemical
sources
Sulfide is toxic to many plants and animals and reacts with
numerous metals
Sulfur-oxidizing chemolithotrophs can oxidize sulfide and
elemental sulfur at oxic/anoxic interfaces

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Catabolic Diversity
Fermentation

Carbon flow

Organic compound

Carbon flow in
respirations

Electron transport/
generation of pmf

Biosynthesis

Aerobic respiration

Chemotrophs

Electron
acceptors

Anaerobic respiration

Chemoorganotrophy

Electron transport/
generation of pmf

Electron
acceptors

Biosynthesis

Aerobic respiration

Anaerobic respiration
Chemolithotrophy
Light

Phototrophs

Photoheterotrophy
Organic
compound

Photoautotrophy

Electron
transport

e
donor

Generation of pmf
and reducing power
Biosynthesis

Biosynthesis

Phototrophy

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Fig. 4.22

Chemoorganotrophy
Chemoorganotrophy
Energy and carbon
source:
organic compound(s)

Fermentation, anaerobic process without


external electron acceptors
Energy generating process:
Substrate-level-phosphorylation (SLP)
Example: Lactic acid fermentation
Catabolic end product: lactic acid (homofermentative)

Chemotrophs

Fermentation
Carbon flow in
respirations

Carbon flow

Organic compound
Electron transport/
generation of pmf

Biosynthesis

Aerobic respiration

Electron
acceptors

Anaerobic respiration
Chemoorganotrophy
Anaerobic respiration, alternative electron
acceptors (not O2)
Energy generating process:
Electron transport phosphorylation (and SLP)
Catabolic end product: CO2
(exception: methanogenesis -> CH4)
Example: Escherichia coli with nitrate

Aerobic respiration, O2 as electron acceptor


Energy generating process:
Electron transport phosphorylation (and SLP)
Catabolic end product: CO2
Example 1: Paracoccus
Example 2: Escherichia coli

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Chemolithotrophy
Energy source: reduced inorganic compound
Electron acceptor, mostly O2
Energy generating process: Electron transport phosphorylation
Catabolic end product: oxidized inorganic compound
Example: Nitrification of Nitro(so)bacteria
Carbon source: CO2 (in most bacteria/archaea)

Electron transport/
generation of pmf
Electron
acceptors

Biosynthesis

Aerobic respiration

Anaerobic respiration

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Phototrophy

Energy source: Light


Energy generating process: Electron transport phosphorylation
Carbon source: CO2 (in most bacteria)

Light

Photoheterotrophy
Organic
compound

Photoautotrophy

Electron
transport

e
donor

Generation of pmf
and reducing power
Biosynthesis

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Biosynthesis